The Cambrian Explosion: Biology’s Big Bang Stephen C. Meyer, Marcus Ross, Paul Nelson, and Paul Chien Q Q Q I. Introduction: Design without a Designer? Both Charles Darwin himself and contemporary neo-Darwinists such as Francisco Ayala, Richard Dawkins, and Richard Lewontin acknowledge that biological organisms appear to have been designed by an intelligence. Yet classical Darwinists and contemporary Darwinists alike have argued that what Francisco Ayala calls the “obvious design” of living things is only apparent. As Ayala, a former president of the American Association for the Advancement of Science, has explained: “The functional design of organ- isms and their features would therefore seem to argue for the existence of a designer. It was Darwin’s greatest accomplishment to show that the di- rective organization of living beings can be explained as the result of a nat- ural process, natural selection, without any need to resort to a Creator or other external agent.”1 According to Darwin and his contemporary followers, the mechanism of natural selection acting on random variation is sufficient to explain the ori- gin of those features of life that once seemed to require explanation by reference to an intelligent or purposeful designer. Thus, according to 323 324 Meyer et al. Darwinists, the design hypothesis now represents an unnecessary and un- parsimonious explanation for the complexity and apparent design of living organisms. On these as well as methodological grounds contemporary biol- ogists have generally excluded the design hypothesis from consideration as an explanation for the origin of biological form. Yet does Darwinism, in either its classical or contemporary versions, fully succeed in explaining the origin of biological form? Can it explain all evi- dence of apparent design? Most biologists now acknowledge that the Dar- winian mechanism of natural selection acting on random variations can explain small-scale microevolutionary changes, such as cyclical variations in the size of the beaks of Galapagos finches or reversible changes in the ex- pression of genes controlling color in English peppered moths.2 But what about the large-scale innovations in the history of life? What about the ori- gin of completely new organs, body plans, and structures—the macroevolu- tionary innovation to which the fossil record attests? Does Darwinism, or neo-Darwinism, or any other strictly materialistic model of evolutionary change explain the origin of the basic body plans or structural “designs” of animal life, without invoking actual (that is, purposive or intelligent) design? In this essay, we will test the claims of neo-Darwinism and two other materialistic models of evolutionary theory: punctuated equilibrium and self-organization. We will do so by assessing how well these theories ex- plain the main features of the Cambrian explosion—a term that refers to the geologically sudden appearance of numerous new animal forms (and their distinctive body plans) 530 million years ago. We shall show that the Cambrian fossil record contradicts the empirical expectations of both neo- Darwinism and punctuated equilibrium in several significant respects. We further show that neither neo-Darwinism’s selection/mutation mechanism nor more recent self-organizational models can account for the origin of the biological information necessary to produce the Cambrian animals and their distinctive body plans. Instead, we will argue that intelligent design explains both the pattern of the fossil record and the origin of new biolog- ical form and information better than the competing models of purposeless and undirected evolutionary change. II. The Cambrian Explosion The term Cambrian explosion describes the geologically sudden appearance of animals in the fossil record during the Cambrian period of geologic time. During this event, at least nineteen, and as many as thirty-five (of forty total), phyla made their first appearance on earth.3 Phyla constitute the The Cambrian Explosion: Biology’s Big Bang 325 highest biological categories in the animal kingdom, with each phylum ex- hibiting a unique architecture, blueprint, or structural body plan. Familiar examples of basic animal body plans are cnidarians (corals and jellyfish), mollusks (squids and shellfish), arthropods (crustaceans, insects, and trilo- bites), echinoderms (sea star and sea urchins), and the chordates, the phy- lum to which all vertebrates including humans belong. The fossils of the Cambrian explosion exhibit several distinctive features. A. Geologically Sudden Appearance and the Absence of Ancestral Precursors or Transitional Intermediates First, as the name implies, the fossils of the Cambrian explosion appear suddenly or abruptly within a very brief period of geologic time (see figures 1 and 2). As recently as 1992, paleontologists thought the Cambrian period began 570 million years ago and ended 510 million years ago, with the Cambrian explosion itself occurring within a 20-to-40-million-year win- dow during the lower Cambrian period. In 1993, radiometric dating of zir- con crystals from formations just above and just below Cambrian strata in Siberia allowed for a precise recalibration of the age of Cambrian strata. Radiometric analyses of these crystals fixed the start of the Cambrian hiopoda niaoida nelida ophora olitha ordata ropoda c o n n y h h Bra Eld A Cte H C Art FIGURE 1. Some of the major animal groups (phyla) that first appeared in the Cambrian explosion. Artistic reconstructions of Cambrian fossils: Brachiopoda by Andrew Johnson, Chordata(Myllokunmingia fengjiaoa) courtesy of D. G. Shu, all other images courtesy of J. Y. Chen. 326 Meyer et al. Present Cambrian period ends: 490 mya Cambrian explosion:530 - 525 mya = 5 million years(One tenth of 1% of Earth’s history) 525 mya 530 mya Cambrian period begins: 543 mya Origin of Earth: 4600 mya FIGURE 2.The Cambrian explosion occurred within a narrow window of geo- logic time. Artistic reconstructions of Cambrian fossils: Brachiopoda by Andrew Johnson, Chordata (Myllokunmingia fengjiaoa) courtesy of D. G. Shu, all other images courtesy of J. Y. Chen. period at 543 million years ago and the beginning of the Cambrian explo- sion itself at 530 million years ago (see figure 2).4 These studies also showed that the Cambrian explosion occurred within an exceedingly nar- row window of geologic time, lasting no more than 5 million years. Geo- logically speaking, 5 million years represents a mere 0.11 percent of Earth’s history. As Chinese paleontologist Jun-Yuan Chen has explained, “com- pared with the 3-plus-billion-year history of life on earth, the period [of the explosion] can be likened to one minute in 24 hours of one day.”5 Yet most of the innovations in the basic architecture of animal forms occurred abruptly within just such a small fraction of the earth’s history during the Cambrian. Due to the suddenness of the appearance of animal life in the Cambrian, the Cambrian explosion has now earned titles such as “The Big Bang of Animal Evolution” (Scientific American), “Evolution’s Big Bang” (Science), and the “Biological Big Bang” (Science News).6 To say that the fauna of the Cambrian period appeared in a geologically sudden manner also implies the absence of clear transitional intermediates connecting the complex Cambrian animals with those simpler living forms found in lower strata. Indeed, in almost all cases, the body plans and struc- tures present in Cambrian period animals have no clear morphological antecedents in earlier strata. Some have argued that perhaps the (or The Cambrian Explosion: Biology’s Big Bang 327 Venelian) fauna hold some hope in this regard, but as we will show below, those hopes now seem unfounded. B. Extensive Morphological Breadth and Representation of Phyla Second, the Cambrian explosion exhibits an extraordinary morphological breadth and representation of the disparate animal phyla. Cambrian rocks dis- play about half (or more) of the basic body plans or architectural designs of the animal kingdom. Representatives of nineteen of the forty known animal phyla definitely make their first appearance in the fossil record during the Cambrian explosion.7 Three phyla appear in the Precambrian. Six animal phyla first appear in the fossil record after the Cambrian period, and twelve more are not represented in the fossil record. Nevertheless, for reasons described below, many paleontologists think that almost all of these additional eighteen phyla may well have originated during the Cambrian explosion. Some authorities even estimate that all animal phyla might have come into existence during the Cambrian explosion. As Valentine, Jablonski, and Erwin argue, “All living phyla may have originated by the end of the [Cambrian] explosion.”8 An especially dramatic feature of the Cambrian explosion is the first appearance of all the invertebrate phyla (and subphyla) with mineralized exoskeletons, including the advanced metazoans such as Mollusca, FIGURE 3. Fossil trilobite found in lower Cambrian strata near Chengjiang, China. Courtesy of J. Y. Chen. Phylum: Arthropoda Subphylum: Trilobitomorpha Genus & species: Eoredlichia intermedia 328 Meyer et al. FIGURE 4.Artistic reconstruction and fossil specimen of an Anomalocarisfound near Chengjiang China. Courtesy of J. Y. Chen. Phylum: Arthropoda Subphylum: Anomalocarida Genus & species: Anomalocaris saron Echinodermata, and Arthropoda. Trilobites (see figure 3), a subphylum of Arthropoda, were highly complex animals whose thoraxes comprised three lobes or sections (a medial axial ring and two lateral pleurae). The bodies of trilobites were covered by a shieldlike, keratinized exoskeleton called a carapace, which covered both the head and thorax of these animals. Like modern arthropods, trilobites grew by shedding their carapaces, and these cast-off carapaces help to account for the abundance of trilobite fossils. The Chengjiang fauna also contains a number of fossils of now-extinct, top-of- the-food-chain predators with exotic names such as Anomalocaris(up to six feet in length; see figure 4), which indicates the presence of a complex food web and a diverse ecological community. Shelled animals leave a far more durable and extensive record than their soft-bodied counterparts. Nevertheless, Cambrian fossil discoveries from both the Burgess shale in the Canadian Rocky Mountains and from The Cambrian Explosion: Biology’s Big Bang 329 FIGURE 5. Artistic reconstruction and fossil specimen of a comb jelly found near Chengjiang, China. Courtesy of J. Y. Chen. Phylum: Ctenophora Genus & species: Maotianoascus octonarius the lower Cambrian Yuanshan Formation near Chengjiang, China, have also shown exquisitely preserved soft-bodied fauna. The Chengjiang fauna even show many excellent examples of well-preserved animals with soft tissue (animals lacking even a keratinized exoskeleton), including mem- bers of phyla such as Cnidaria, Ctenophora (see figure 5), Annelida, Ony- cophora, Phoronida (see figure 6), and Priapulida. Burgess Shale fossils from the middle Cambrian (515 million years ago) confirm that many of these Cambrian organisms were long-lived and geographically widespread. The lower Cambrian sediments near Chengjiang have preserved fossils of such excellent quality that soft tissues and organs, such as eyes, intes- tines, stomachs, digestive glands, sensory organs, epidermis, bristles, mouths, and nerves, can be observed in detail. Even fossilized embryos of sponges are present in the Precambrian strata near Chengjiang.9 Cam- brian-level strata show the soft body parts of jellyfish–like organisms (known as Eldonia; see figure 7), such as radiating water canals and nerve rings. These fossils even include the gut contents of several different kinds of animals and undigested food residue in their stools.10 330 Meyer et al. FIGURE 6.Artistic reconstruction and fossil specimen of a phoronid found near Chengjiang, China. Courtesy of J. Y. Chen. Phylum: Phoronida Genus & species: Iotuba chengjiangensis The Chengjiang fauna also confirms the presence of animals from the phylum Chordata. Yunnanozoon lividum is a fusiform eel-shaped animal with, among other features, a digestive tract, branchial arches, and a large notochord. Yunnanozoon has been interpreted as a primitive chordate.11 Two possible cephalochordates have been identified from Cambrian de- posits in both China and Canada: Paleontologists have found a single spec- imen of a possible cephalochordate, Cathaymyrus,from the lower Cambrian Qiongzhusi Formation near Chengjiang.12 Additionally, the cephalochor- date Pikaia is known from the middle Cambrian Burgess Shale.13 More important, several recent discoveries from China have surprised (if not shocked) the paleontological community regarding early chordates. Chen, Huang, and Li have recently reported the discovery of a sophisti- cated craniate-like chordate called Haikouella lancelotafrom the lower Cam- brian Maotianshan Shale in China. According to Chen and his colleagues, Haikouella has many of the same features of the Yunnanozoon lividum, as well as several additional anatomical features including a “heart, ventral and dorsal aorta, an anterior branchial arterial, gill filaments, a caudal pro- jection, a neural cord with a relatively large brain, a head with possible The Cambrian Explosion: Biology’s Big Bang 331 FIGURE 7.Artistic reconstruction and fossil specimen of an Eldoniafound near Chengjiang, China. Courtesy of J. Y. Chen. Phylum: Eldoniaoida(contested designation) Genus & species: Eldonia eumorpha lateral eyes, and a ventrally situated buccal cavity with short tentacles.”14 Also, D. G. Shu, Simon Conway Morris, and several Chinese colleagues have made a dramatic find of two small fish fossils, Myllokunmingia fengjiaoa and Haikouichthys ercaicunensis (see figures 8 and 9) suggesting a much ear- lier appearance for vertebrates than previously thought. Both of these taxa are jawless fish (agnathans) and are considered by Shu and his colleagues to be rather closely allied to lampreys.15 Lastly, a new paper by Shu and others reports the first convincing spec- imen of a urochordate (tunicate) from the Cambrian.16This specimen, Che- ungkongella ancestralis, is likewise found from the early Cambrian shales (Qiongzhusi Formation) near Chengjiang. Remarkably, these recent finds now demonstrate that not only was the phylum Chordata present in the Cambrian but also that each one of the chordate subphyla (Cephalochor- data, Craniata, and Urochordata) were present. Moreover, each of these chordate taxa displayed the morphological characteristics that place them securely within their respective subphyla. Indeed, many phyla such as arthropods, mollusks, and chordates include morphologically disparate subphyla that many paleontologists regard as separate body plans. If subphyla are included in the count of animal body plans, then at least thirty-two and possibly as many as forty-eight of fifty-six total body plans (57.1 to 85.7 percent) first appear on earth during the Cambrian explosion. Thus, depending on how one evaluates the data at either the phyla or FIGURE 8.Artistic reconstruction and fossil specimen of a fish (similar to mod- ern hagfish) found near Haikou, China. Courtesy of D. G. Shu. Phylum: Chordata Subphylum: Vertebrata Genus & species: Myllokunmingia fengjiaoa FIGURE 9.Artistic reconstruction and fossil specimen of a fish (similar to mod- ern lamprey) found near Haikou, China. Courtesy of D. G. Shu. Phylum: Chordata Subphylum: Vertebrata Genus & species: Haikouichthys ercaicunensis